Cerebral Cortex April 2010;20:912--928
doi:10.1093/cercor/bhp153
Advance Access publication August 13, 2009
Tracking Lexical Access in SpeechProduction: ElectrophysiologicalCorrelates of Word Frequency andCognate Effects
Kristof Strijkers1,2, Albert Costa2 and Guillaume Thierry3
1Universitat de Barcelona, Department of Psicologia Basica,
GRNC, 08035 Barcelona, Cataluna, Spain, 2Universitat Pompeu
Fabra, Dept. de Tecnologia, ICREA, 08018 Barcelona, Cataluna,
Spain and 3Bangor University, School of Psychology, ESRC
Centre for Research on Bilingualism in Theory and Practice,
Bangor, LL57 2DG Wales, UK
The present study establishes an electrophysiological index oflexical access in speech production by exploring the locus of thefrequency and cognate effects during overt naming. We conducted2 event-related potential (ERP) studies with 16 Spanish--Catalanbilinguals performing a picture naming task in Spanish (L1) and 16Catalan--Spanish bilinguals performing a picture naming task inSpanish (L2). Behavioral results showed a clear frequency effectand an interaction between frequency and cognate status. The ERPelicited during the production of high-frequency words divergedfrom the low-frequency ERP between 150 and 200 ms post-targetpresentation and kept diverging until voice onset. The same resultswere obtained when comparing cognate and noncognate con-ditions. Positive correlations were observed between naminglatencies and mean amplitude of the P2 component following thedivergence, for both the lexical frequency and the cognate effects.We conclude that lexical access during picture naming beginsapproximately 180 ms after picture presentation. Furthermore,these results offer direct electrophysiological evidence for an earlyinfluence of frequency and cognate status in speech production.The theoretical implications of these findings for models of speechproduction are discussed.
Keywords: ERP, language production, lexical activation, time course
Introduction
The ease with which we produce speech may lead us to think
that the cognitive and brain mechanisms put at play in this skill
are rather simple. However, speech production is a complex
process which entails the orchestration of many processes that
unfold over time (e.g., Dell 1986; Caramazza 1997; Levelt et al.
1999). In recent years, the amount of psycholinguistic
experimental research exploring these processes has in-
creased, leading to more detailed models of speech production.
However, the same cannot be said regarding the investigation
of the time course of the neural events underpinning these
processes. The present article aims at helping to fill this gap by
exploring the electrophysiological correlates of 2 robust
psycholinguistic phenomena in picture naming; the frequency
and the cognate effect.
Cognitive models of single word production assume that the
translation of our communicative goal into speech occurs at
various levels of representation. Speaking probably involves at
least 1) the retrieval of conceptual information, 2) the selection
of the words corresponding to the intended message, 3) the
phonological encoding of the selected words, and 4) the
retrieval of the articulatory plans (e.g., Dell 1986; Caramazza
1997; Levelt et al. 1999). Although there is still a debate
regarding how these processes unfold over time, researchers
generally agree on the existence of some sequentiality. That is,
concepts are retrieved before words are selected and these in
turn are selected before their corresponding phonemes are
retrieved. Given this general functional architecture, it is
relevant to describe not only the neural structures implicated
in the representation of different types of information but also
the time course of their involvement.
The time course of lexical access in speech production has
been studied using a variety of chronometric tasks (e.g.,
Schriefers et al. 1990; Dell and O’Seaghdha 1991; Wheeldon
and Levelt 1995). These studies have provided evidence for the
hypothesis of sequentiality in speech production by showing
that a word’s conceptual/semantic and syntactic properties are
retrieved before its phonological form becomes available.
Recently, event-related potential (ERP) studies have supported
such a sequence of processing in speech production (e.g., Van
Turennout et al. 1997, 1998; Schmitt et al. 2000; Jescheniak et al.
2002; Rodriguez-Fornells et al. 2002; Schiller et al. 2003a). These
studies used the Lateralized Readiness Potential (LRP; e.g., Coles,
1989; Miller et al. 1992) and the N200 (a component supposed
to reflect response inhibition; e.g., Pfefferbaum et al. 1985; Kok
1986; Eimer 1993) in linguistic go/no-go tasks in order to obtain
precise measurements of the temporal distance between
different stages of speech production. LRP and N200 data
indicate that conceptual activation unfolds during the first 150
ms of processing (e.g., Thorpe et al. 1996; see also Johnson and
Olshausen 2005; Hauk et al. 2007), lexico-semantic information
is processed 90--120 ms before phonological information (e.g.,
Van Turennout et al. 1997; Schmitt et al. 2000), and syntactic
information is retrieved approximately 40 ms before phonolog-
ical information (e.g., Van Turennout et al. 1998).
In an interesting meta-analysis, Indefrey and Levelt (2004)
integrated results of such chronometric ERP studies with the
time course of neural activation revealed by magnetoencepha-
lography (MEG) in overt picture naming tasks (Salmelin et al.
1994; Levelt et al. 1998; Maess et al. 2002). According to this
meta-analysis and relative to picture onset (time 0), 1) lexical
access (understood as lemma retrieval) is estimated to start as
early as 150 ms and reach completion at around 275 ms, 2)
phonological encoding is thought to take place between 275
and 400 ms; and 3) syllabification is estimated to unfold
between 400 and 600 ms.
However, the chronometric interpretation proposed by
Indefrey and Levelt’s (2004) meta-analysis may be affected by
the fact that the tasks that participants performed in the ERP
studies cited above differed markedly from those used in the
MEG studies considered for comparison. In order to avoid
effects of speech-related motor artifacts (e.g., Wohlert 1993;
Masaki et al. 2001) none of the ERP experiments actually
� The Author 2009. Published by Oxford University Press. All rights reserved.
For permissions, please e-mail: [email protected]
by guest on Septem
ber 30, 2011cercor.oxfordjournals.org
Dow
nloaded from
involved speech production, but rather consisted of button
press responses. Furthermore, the tasks were complex and
difficult (go/no-go decision, combined with left/right button
press decision, followed by production of a pronoun sentence
comprising the stimuli), and implied meta-linguistic judgments
by the participants (e.g., ‘‘if the word refers to an animal, then
press the right button if the animal name starts with the letter
b or press the left button if it starts with the letter s’’). It cannot
be excluded that such conditions may have triggered additional
cognitive processes affecting the timing of the key processes
involved in natural speech production. Also, the time course
proposed by models of picture naming is at odds with some
time course estimates derived from picture processing and
word recognition studies. Whereas some authors argue that
during picture processing the brain engages in semantic
analysis already before 150 ms after stimulus onset (e.g.,
Thorpe et al. 1996; Johnson and Olshausen 2005; Hauk et al.
2007), others dispute the existence of such early differences
(e.g., Holcomb and McPherson 1994; Kiefer 2001; Eddy et al.
2006; Sitnikova et al. 2006). Similarly, some ERP studies of visual
word recognition converge in showing that lexical processing
starts within 200 ms of picture onset (e.g., Sereno et al. 1998;
Hauk and Pulvermuller 2004; Hauk et al. 2006, 2009) but other
studies using the masked priming paradigm showed that it
takes at least 250--300 ms to start retrieving lexical information
(e.g., Holcomb and Grainger 2006; Grainger and Holcomb
2009). Here we sought to obtain new evidence regarding the
time course of lexical access in natural speech production by
asking high proficient bilinguals to simply name pictures
overtly while recording ERPs.
As hinted above, recording electroencephalography (EEG)
signals during overt speech is methodologically disputable, due
to articulation-related artefacts. Indeed, activation of the mouth
and face musculature produces electrical potentials larger than
brain-generated signals by a factor of 10--100. These large
motor and motor preparation potentials, well beyond the EEG
amplitudes collected in silent response tasks such as button-
presses, could potentially overshadow the cognitive brain
activity at interest. However, one ERP study, directly comparing
overt versus covert speech (Eulitz et al. 2000), and several MEG
studies of overt picture naming (e.g., Levelt et al. 1998; Salmelin
et al. 1994; Maess et al. 2002) have shown that reliable
measures of brain activity can be taken at least until 400 ms
after picture onset. As far as we are interested in early phases of
picture naming, this simple paradigm should offer insight into
the time course of lexical retrieval.
In order to trigger ERP differences during lexical access in
speech production, we chose to manipulate word frequency as
an independent variable. There is ample evidence that word
frequency affects the speed and accuracy with which picture
naming is performed: pictures with high-frequency names tend
to be named faster and more accurately by normal and brain-
damaged speakers than pictures with low-frequency names (e.g.,
Oldfield and Wingfield 1968; Dell 1990; Jescheniak and Levelt
1994; Navarrete et al. 2006; Almeida et al. 2007; Kittridge et al.
2007). These results suggest that lexical frequency influences or
even determines the speed of lexical access (for a similar
approach in language comprehension see e.g., King and Kutas
1998; Sereno et al. 1998; Hauk and Pulvermuller 2004).
However, 2 issues must be kept in mind: 1) lexical frequency
correlates with conceptual variables such as imageability and
familiarity and 2) the stage at which word frequency exerts its
effects remains debated. These 2 considerations pose difficulties
for the interpretation of ERP differences driven by lexical
frequency.
One way to circumvent issue (1) is to manipulate another
independent variable that exerts a reliable and consistent effect
in picture naming but that is not confounded by conceptual
properties. In the present study we tested bilingual individuals
and we manipulated not only lexical frequency but also the
cognate status of target picture names. The cognate effect
refers to the observation of faster naming latencies in bilingual
speakers for pictures whose translations are phonologically
similar across languages (e.g., the Spanish--English pair ‘‘gui-
tarra’’— guitar) as compared with pictures with phonologically
dissimilar translations (e.g., the Spanish--English pair ‘‘perro’’—
dog; e.g., Costa et al. 2000, 2005; Christoffels et al. 2007).
Critically, the cognate status of a word depends on its
phonology (formal similarity across translations) and is
therefore unrelated to conceptual variables of the sort de-
scribed above (see description of stimuli and appendix B).
(There has been one proposal arguing that cognate status
might influence conceptual processing; Van Hell and De Groot
1998. Cognates should have larger conceptual overlap com-
pared with noncognates. However, such an account has
difficulties in explaining the performance of brain-damaged
speakers or certain tip-of-the-tongue states and seems to have
problems with theoretical logic [for a clear overview see Costa
et al. 2005]. Furthermore, results from Van Hell and De Groot’s
(1998) study are not that clear for concrete cognates [the type
of stimuli used here] and are based on the sole assumption that
word association does not involve lexical processes. Finally, the
absence of any significant differences between cognate and
noncognate words for the conceptual ratings of the stimuli
used in the present study is at odds [at least for concrete
cognates in high proficient bilinguals] with a conceptual
account.). In a recent ERP study of overt naming involving
cognates, Christoffels et al. (2007) found a significant negative
enhancement around 300 ms after stimulus presentation for
cognate ERPs compared with noncognate ERPs, which was
interpreted in support of the phonological origin of the
cognate effect.
Issue (2), that is, the locus of the frequency effect in speech
production, is more complex to address. According to some
researchers, word frequency affects the retrieval of lexical
nodes from the lexicon (e.g., Caramazza et al. 2001; Navarrete
et al. 2006; Almeida et al. 2007), whereas others argue that
frequency only affects the retrieval of phonological information
during production (e.g., Jescheniak and Levelt 1994; Jescheniak
et al. 2003). Our study can only be informative regarding the
time course of lexical access if indeed word frequency
influences the retrieval of lexical representations and not just
the retrieval of phonological segments. However, we believe
this issue to be an empirical one. Importantly, recent
behavioral, neuroimage and patient studies all provide clear
evidence supporting the notion that frequency affects both
lexical and phonological processing in speech production (e.g.,
Navarrete et al. 2006; Graves et al. 2007; Kittridge et al. 2007;
Knobel et al. 2008). Thus, according to this new evidence, it is
reasonable to expect early effects for frequency (150--200 ms,
e.g., Indefrey and Levelt 2004).
To summarize, our main goal was to index the onset of
lexical access in speech production by comparing ERP differ-
ences elicited by word frequency and cognate status in highly
Cerebral Cortex April 2010, V 20 N 4 913
by guest on Septem
ber 30, 2011cercor.oxfordjournals.org
Dow
nloaded from
proficient Spanish--Catalan speakers naming pictures in L1
(Experiment 1) and highly proficient Catalan--Spanish speakers
naming pictures in L2 (Experiment 2). We hypothesized that
the point in time where the frequency and cognate effects
induced significant ERP differences (i.e., the point in time
where the ERPs start to diverge) would provide new insights
regarding the onset of lexical access. We referred to Indefrey
and Levelt’s (2004) study to predict the time windows in which
the effects of word frequency and cognate status should be
expected. In the case of word frequency, we predicted to find
an early lexical modulation between 150 and 200 ms. For the
cognate effect, based on the ERP study of Christoffels et al.
(2007), we predicted to find a slightly later modulation (after
275 ms).
Note that our predictions are solely based on temporal
information and not bound to the modulation of specific ERP
components. The advantage of such a design is that it will
strongly simplify the interpretation of results. It is noteworthy
that not many studies have involved the recording of ERPs
during overt naming (e.g., Christoffels et al. 2007; Ganushchak
and Schiller 2008; Koester and Schiller 2008; Verhoef et al.
2009). However, these studies have investigated bilingual
language control, error monitoring and morphological priming
in picture--word interference, respectively; therefore the
patterns of brain activity elicited in the current study, that is,
in the context of simple picture naming, maybe quite different.
Experiment 1: Highly Proficient Spanish--Catalan Bilinguals NamingPictures in L1
Method
Participants
Eighteen participants took part in the experiment. All were
highly proficient early Spanish--Catalan bilinguals (see Appen-
dix A) exposed almost exclusively to Spanish during their first
3--4 years of life and reported to have a preference or
dominance for that language. Participants were students at
the University of Barcelona (ages 18--25) and received course
credits or monetary compensation for their participation. All
were right-handed, had normal or corrected-to-normal vision
and did not suffer from any neurological or motor problems.
Two of the participants had to be removed from the analyses:
one due to an unacceptable number of EEG artefacts, and
another due to technical problems during EEG recording.
Statistical analyses are thus based on 16 individual data sets.
Materials
Sixty-four line-drawings of familiar objects corresponding to
Spanish words were selected, spanning a wide range of semantic
categories (e.g., body parts, buildings, animals, furniture; see
Fig. 1). Two independent variables were manipulated orthogo-
nally: cognate status and lexical frequency of the picture names.
This design therefore entailed 4 experimental conditions involv-
ing 16 pictures each (see Appendix B). Pictures with high-
frequency names were at least 10 times more frequent than
pictures with low-frequency names (mean lemma frequency
[LEXESP; Sebastian-Galles et al. 2000]: low frequency: 3.9; high
frequency: 52). The mean lexical frequency of the picture
names in the cognate and noncognate sets were similar (non-
cognates: 27.8; cognates: 32.2). The cognate words shared on
average 4 phonological segments with their translation equiv-
alents (range = 2--8). Almost all cognates (27 out of 32) shared at
least the whole first syllable with their corresponding trans-
lations, and all of them shared at least the first phoneme. There
was no obvious phonological or orthographic overlap between
noncognates. None shared their first phoneme and only 7 out of
32 shared the first vowel appearing in a word. Words in the 4
conditions were also matched for length (mean syllable length:
low-frequency noncognates: 2.7; low-frequency cognates: 2.6;
high-frequency noncognates: 2.4; high-frequency cognates: 2.2).
Physical variance within each of the 4 stimulus sets was
evaluated using interstimulus pixel-wise correlations (Thierry
et al. 2007), and no difference was found between experimental
conditions (Fig. 1). In addition, 40 students who did not par-
ticipate in the study rated the complexity of the stimuli using
a Spanish adaptation of the methods described in Snodgrass and
Vanderwant’s (1980) study. We found no differences between
any of the 4 stimulus sets taken 2-by-2 (P > 0.2). Finally, all items
used in the experiment were rated on 3 conceptual variables
(familiarity, typicality, and imageability) by 120 students not
tested in the ERP experiment, using a Spanish adaptation of the
Snodgrass and Vanderwart (1980) procedure. Low- and high-
frequency items only differed significantly (P < 0.001) for the
familiarity ratings but not on typicality or imageability (P > 0.7).
Critically, there were no differences between cognates and
noncognates for any of the 3 conceptual properties (P > 0.5).
Finally, to increase experimental power, each picture was
presented once in each of 6 separate blocks (which makes 64
stimuli per block) with order of the presentation within blocks
randomized for each participant.
Procedure
Participants were tested individually in a soundproof room.
Instructions were administered in Spanish. Participants were
Figure 1. Exemplars of picture stimuli used in the 4 experimental conditions andtheir names in Spanish and Catalan. The images on the right hand side and thebottom depict the standard deviation of each individual picture in a series from themean image in that series. Black shows no difference with the mean, white showsmaximal difference from the mean. There is no differential pattern emerging fromthese standard deviation images, which indicates no systematic bias in interstimulusvariability between experimental conditions.
914 The Time Course of Lexical Access in Speech Production d Strijkers et al.
by guest on Septem
ber 30, 2011cercor.oxfordjournals.org
Dow
nloaded from
asked to name each picture presented in Spanish as fast and as
accurately as possible. Before the experiment started, partic-
ipants were familiarized with the pictures and they were given
feedback (correct picture name) when they made a non
response or an incorrect response (23%). Each experimental
trial had the following structure: 1) a blank interval of 1000 ms
was shown at the centre of a computer screen; 2) a picture was
displayed at the centre of the screen until a response was given
or for a maximum of 1500 ms; 3) a blank intertrial interval of
1000 ms. An asterisk was presented for 500 ms before the first
trial and after the last trial to signal the beginning and end of
each block. Blocks were separated by a 30-s pause. Response
latencies were measured from the onset of the stimulus by
means of a voice key. Stimulus presentation was controlled by
an adaptation of the EXPE Program (Pallier et al. 1997). The
entire experimental session lasted approximately 25 min. At
the end of the experimental session, participants were asked to
fill in a questionnaire regarding language use and proficiency in
their 2 languages.
EEG Procedure
The EEG was continuously recorded from 31 scalp locations
(Fp1, Fpz, Fp2, F3, Fz, F4, F7, F8, FC1, FC2, FC5, FC6, C3, Cz, C4,
CP1, CP2, CP5, CP6, P3 , Pz, P4, PO1, PO2, T3, T4, T5, T6, O1,
Oz, O2) using tin electrodes embedded in an elastic cap. Five
additional electrodes were placed on the participants’ head.
Two bipolar electrodes were placed next to and beneath the
left eye (EOGH and EOGV) to register eye movements; 2 other
electrodes were placed on the participants’ right and left
mastoid (A1 and A2) for on-line referencing and a final
electrode was placed on the participants’ nose as off-line
reference channel. The EEG was continuously recorded and
digitized at 250 Hz. Impedances were reduced to 3 kOhms or
less prior to the beginning of recording. Before segmentation,
the EEG was processed through a low-pass filter with a cut-off
frequency of 20 Hz and a high-pass filter of 0.03 Hz. The EEG
was then segmented into 750-ms-long epochs starting 200 ms
prior to stimulus onset. We chose to segment the EEG only up
to 550 ms after stimulus onset to avoid speech contamination
(e.g., Wohlert 1993; Masaki et al. 2001). Just as in Christoffels
et al. (2007), Verhoef et al. (2009), and Koester and Schiller
(2008) we assumed that analyzing the ERPs before the actual
response would result in artifact-free ERPs. In contrast to those
studies, however, we chose to segment in such a way that 1)
fast responses could not induce motor artifacts and 2) latency
jitter by strong EMG activity could be avoided in the averaging.
For the naming latencies, the fastest average response was
650 ms. We segmented the EEG up to 550 ms (100 ms less than
the fastest average response, to ensure that we could include as
many epochs as possible) and removed prior to averaging all
epochs where the response was faster than 550 ms. The
segmented EEG underwent Gratton and Coles (1989) ocular
correction and artifact rejection where trials with an amplitude
voltage over 100 lV or a change in amplitude between adjacent
samples of more than 200 lV were dismissed. Also trials where
participants’ responses were incorrect or absent and trials
containing other motor artifacts were rejected from the dataset
before averaging. The 750-ms epochs were then averaged in
reference to the 200-ms prestimulus baseline. In total, ERP
analyses were based on an average of 162 segments (SD = 21)
per condition taken 2-by-2 (e.g., low-frequency noncognates +low-frequency cognates for the low-frequency condition) per
subject (low frequency: 164, high frequency: 160, noncognate:
164, cognate: 160).
Analyses
Error Analysis
Four types of responses were scored as errors: 1) production of
incorrect names; 2) verbal disfluencies (stuttering, utterance
repairs, production of nonverbal sounds that triggered the
voice key); 3) recording failures; 4) errors in which the
bilingual participants named the picture in Catalan. This gave
a total of 1.2% erroneous responses. Furthermore, outliers (i.e.,
responses exceeding 3 standard deviations from the partic-
ipant’s mean, 2.7%) and responses faster than 550 ms (10.6%)
were excluded from all analyses. For the ERP analysis another
4.7% of the data were excluded due to artefacts. Given the very
low amount of erroneous responses (1.2%) we decided not to
run a statistical error analysis.
Behavioral Analyses
Separate subject (F1) and item (F2) analyses were carried out
examining 3 independent variables: Cognate Status (cognates
vs. noncognates), Frequency Status (low frequency vs. high
frequency) and Block (6 repetitions).
ERP Analysis
The main goal was to identify the latency at which the ERPs of
the 2 contrasts of interest (low-frequency vs. high-frequency
ERPs and cognate vs. noncognate ERPs) started to diverge
significantly from one another. We adopted a method sug-
gested by Guthrie and Buchwald (1991; see also Thierry et al.
1998, 2003). ERPs were compared between conditions at each
electrode by running 2-tailed paired t-tests at every sampling
point (4 ms) starting from target presentation (0 ms) until at
least a sequence of 12 consecutive t-test samples exceeded the
0.05 significance level. We also estimated the splitting point,
that is, the point in time where ERPs started to diverge in each
individual subject, in order to perform a correlation analysis
between this splitting point and the mean naming latencies of
each subject.
Second, 3 time window analyses were conducted to explore
possible interactions between frequency and cognate status,
and to explore the effect of repetition in the ERPs: 1) a 2 3 2 3
9 repeated measures ANOVA was conducted with Frequency
Status (low frequency vs. high frequency), Cognate Status
(noncognate vs. cognate), and Electrode (9 electrode clusters;
see below) as independent variables; 2) a 2 3 6 3 9 ANOVA was
conducted with Frequency Status, Block (6 blocks) and
Electrode as independent variables; 3) a 2 3 6 3 9 ANOVA
was conducted with Cognate Status, Block, and Electrode as
independent variables. Five time windows were selected, based
upon visual inspection of the Grand Averages: 1) 0--80 ms, 2)
80--160 ms, 3) 160--240 ms (P2), 4) 240--320 ms (N3), and 5)
320--420 ms (P3). Note that P2, N3 and P3 are used as
descriptive labels here. Electrodes were clustered in 9 groups
as follows: Left frontal (Lfr): F7,F3,FC5; Fronto-central (Fc):
Fz,FC1,FC2,Cz; Right frontal (Rfr): F8,F4,FC6; Left Central (Lc):
T3,C3,CP5; Centro-Parietal (Cpar): CP1,CP2,Pz; Right central
(Rc): T4,C4,CP6; Left Parietal (Lpar): T5,P3,PO1; Occipital (Oc):
O1,Oz,O2; Right Parietal (Rpar): T6,P4,PO2).
Thirdly, because the method presented here is new for
studying speech production, correlation analyses were
Cerebral Cortex April 2010, V 20 N 4 915
by guest on Septem
ber 30, 2011cercor.oxfordjournals.org
Dow
nloaded from
performed to help us understand the relationship between the
different phases of information processing indexed by ERPs and
the relationship of these phases with behavioral results. On the
one hand, correlation analyses were performed on the in-
dividual splitting point in the ERPs with the individual mean
naming latencies for the high frequency and cognate condition,
because these are the measures likely to show whether or not
the spitting point indexes the onset of lexical access. On the
other hand, correlation analyses were performed on the peak
latencies of the P2, N3, and P3. Peak latencies were measured at
the electrode of maximum amplitude for each component in
each subject (Picton et al. 2000). Finally, a correlation analysis
was performed on individual differences in naming latencies
(the frequency effect and the cognate effect in each subject)
and the individual mean amplitude differences for each ERP
processing phase in each subject. All correlation analyses were
conducted making use of the same electrode clusters as for the
time window analyses.
Behavioral Results
In the analysis of the naming latencies the main effects of Block
(F1(5,75) = 8.9, MSE = 4867.9, P < 0.001; F2(5,300) = 14.7,
MSE = 2139.9, P < 0.001) and Frequency Status (F1(1,15) =47.9, MSE = 3564.1, P < 0.001; F2(1,60) = 8.5, MSE = 19,890.2,
P = 0.005) were significant, and a marginally significant effect of
Cognate Status was found (F1(1,15) = 42.1, MSE = 1603.9, P <
0.001; F2(1,60) = 3.7, MSE = 19,890.2, P = 0.060). Participants
named pictures with high-frequency names faster than pictures
with low-frequency names, and pictures with cognate names
faster than pictures with noncognate names (see Table 1). The
interaction between Frequency Status and Cognate Status was
significant only in the analysis by subjects (F1(1,15) = 8.0, MSE =2484.8, P = 0.013; F2(1,60) = 1.4, MSE = 19,890.2, P = 0.250).
That is, the cognate effect was larger for high-frequency words
(41 ms) than for low-frequency words (12 ms). Finally, there
was also a significant interaction between Cognate Status and
Block in the subject analysis (F1(5,75) = 3.6, MSE = 1088.5, P =0.014; F2 < 1). However as can be seen in Figure 2 this
interaction was mostly driven by the much smaller cognate
effect in the second block compared with the other blocks.
None of the other interactions were significant (P > 0.1; see
Fig. 2).
ERP Results
Onset Latency Analyses
ERPs displayed a typical P1--N1--P2 peak sequence classically
observed for visual stimulus presentation. t-Tests at each
sampling rate indicated that high-frequency ERPs started to
diverge significantly from low-frequency ERPs 172 ms after
picture onset (see Figs 3a and 4a). As can be seen in Figure 4a,
the distribution of electrodes showing a significant effect at
this time point was particularly left-lateralized (however 4 ms
later, almost all electrodes showed significant differences). The
averaged splitting point computed from individual splitting
point estimates was 167 ms, that is, almost within one time
sampling unit of the splitting time measured in the grand-
averages. Cognate ERPs started to diverge significantly from
noncognate ERPs 200 ms after picture onset (see Figs 3b and
4b). The distribution of electrodes displaying significant differ-
ences at this time point was more right-lateralized (but also
here at the next time point practically all electrodes showed
significant differences). Again the grand-average splitting time
of 200 ms closely resembled the averaged individual splitting
point (192 ms). The difference in onset between the splitting
point of the frequency effect and the splitting point of the
cognate effect was significant (measured by individual splitting
points; P = 0.05).
Time Window Analyses
Early time windows (0--80 ms and 80--160 ms). The only
effect found for all 3 ANOVAs conducted in the early time
windows was a small trend for Block between 80 and 160 ms
(F(5,75) = 2.2, MSE = 20,943.4, P = 0.098). In comparison to the
first repetition, all subsequent repetition ERPs seemed to be
more positive going. However, independent ANOVAs of the
possible contrasts with correction for multiple comparisons
showed no significant effects (P > 0.150). None of the other
main effects or interactions were significant (P > 0.175).
Late time windows (160--240 ms (P2), 240--320 ms (N3),
and 320--420 ms (P3). In all 3 time windows significant main
effects were present for Frequency Status (P2: F(1,15) = 8.9,
MSE = 13.6, P = 0.009; N3: F(1,15) = 33.1, MSE = 31.1, P <
0.001; P3: F(1,15) = 39.1, MSE = 18.8, P < 0.001) and Cognate
Status (P2: F(1,15) = 10.6, MSE = 12.1, P = 0.005; N3: F(1,15) =49.9, MSE = 25, P < 0.001; P3: F(1,15) = 28.1, MSE = 22.4, P <
0.001). ERPs in the high-frequency condition were signifi-
cantly more negative than those elicited in the low-frequency
condition and ERPs in the cognate condition were signifi-
cantly more negative than those elicited in the noncognate
condition. The distribution of this effect emerged at posterior
Table 1Mean naming latencies (ms) in the 4 experimental conditions for Experiment 1 and Experiment 2
Low-frequencynoncognates(ms)
Low-frequencycognates(ms)
High-frequencynoncognates(ms)
High-frequencycognates(ms)
Experiment 1(naming in L1)
730 718 702 661
Experiment 2(naming in L2)
764 742 737 694
620
660
700
740
780
820
1 2 3 4 5 6Blocks
Nam
ing
Late
ncie
s (m
s)
LFNC
LFC
HFNC
HFC
Figure 2. Mean naming latencies in Experiment 1 (LFNC 5 low-frequencynoncognate; LFC 5 low-frequency cognate; HFNC 5 high-frequency noncognate;HFC 5 high-frequency cognate) over repetitions.
916 The Time Course of Lexical Access in Speech Production d Strijkers et al.
by guest on Septem
ber 30, 2011cercor.oxfordjournals.org
Dow
nloaded from
sites of the scalp, but expanded rapidly over large parts of the
entire scalp (see Fig. 5). The interaction between Frequency
Status and Cognate Status was significant, but only for the 2
latest time windows (P2: F < 1; N3: F(1,15) = 6.9, MSE = 14, P =0.019; P3: F(1,15) = 16.6, MSE = 9.4, P = 0.001). Starting around
240-ms poststimulus presentation, amplitude differences
between cognate and noncognate ERPs were larger in the
high than in the low-frequency condition. There was neither
a main effect of Block nor interactions of Block with
Frequency Status and/or Cognate Status in any of the 3 time
windows (P > 0.45).
Correlation Analyses
Pearsons product--moment correlation analyses showed no
significant correlation between the individual splitting point
of the low-frequency versus high-frequency ERPs with the
individual mean naming latencies of the high-frequency
condition, and no significant correlation between the in-
dividual splitting point of the noncognate versus cognate ERPs
with the individual mean naming latencies of the cognate
condition (P > 0.6). The correlations between the individual
peak latencies of the P2, N3, and P3 were not significant
either (P > 0.2).
Fz F4
FC1 FC2
CzC3 C4
CP1 CP2
PzP3 P4
PO2PO1
O2OzO1
-5 [µV]
200 400 550
P2
P3
N3
High Frequency Low Frequency
F4Fz
FC2
Cz
CP2
Pz
PO2
O2Oz
Cognates Non-Cognates
(a)
(b)
C4
P4
-5 [µV]
200 400 550
F3
FC1
C3
CP1
P3
PO1
O1
F3
Figure 3. (a) Low-frequency ERPs compared with high-frequency ERPs in Experiment 1 at anterior, central, and posterior scalp locations. Low-frequency ERPs are representedby a dotted line and high-frequency ERPs by a full line. Negativity is plotted upwards. (b) Noncognate ERPs compared with cognate ERPs of Experiment 1 at anterior, central, andposterior scalp locations. Noncognate ERPs are represented by a dotted line and Cognate ERPs by a full line. Negativity is plotted upwards.
Cerebral Cortex April 2010, V 20 N 4 917
by guest on Septem
ber 30, 2011cercor.oxfordjournals.org
Dow
nloaded from
Significant positive Pearson product--moment correlations
were found in the P2 range between the difference in mean
amplitude between high and low-frequency ERPs and the high-/
low-frequency naming latency difference over left-parietal (R =0.531, P = 0.034) and right parietal electrodes (R = 0.511, P =0.043), and a trend towards a positive correlation for the P2 at
the right-central electrodes (R = 0.455, P = 0.077): The larger the
difference in naming latencies between high- and low-frequency
words, the larger the difference in P2 mean amplitude between
high- and low-frequency conditions.
In the N3 range, only a small trend was found in the same
direction over left (R = 0.477, P = 0.082) and right parietal (R =0.429, P = 0.098) electrode clusters. Finally, in the P3 range,
there were no significant correlations between the frequency
effect in the naming latencies and the mean amplitude
differences in the ERPs (P > 0.250).
Analyses of the cognate effect revealed a remarkably similar
pattern of correlations: Trends towards positive correlations
were found for the cognate effect in the P2 range over left
parietal (R = 0.465, P = 0.070) and right parietal (R = 0.436, P =0.091) electrode clusters. The trend for correlation found in
the P2 range disappeared in the N3 (P > 0.190) and P3 (P >
0.475) ranges. Both the frequency effect and the cognate effect
seemed to relate mostly to the early P2.
Discussion
As expected pictures with high-frequency names yielded faster
naming latencies than those with low-frequency names, and
cognate picture names were produced faster as compared
with noncognate picture names. Interestingly, high-frequency
ERPs started to diverge significantly from low-frequency ERPs
172-ms post-target presentation, with high-frequency picture
names eliciting greater negativities than low-frequency names.
Similar results were found for the cognates: cognate ERPs
started to diverge significantly from the noncognate ERPs
200-ms postpicture presentation with more negativity in the
cognate than in the noncognate condition. For both the fre-
quency and the cognate manipulations, these differences
remained from the moment of the split until the end of the
epoch. The absence of peak latency correlations between the
components that were significantly modulated (P2, N3, P3)
suggests that these components have different functional
underpinnings and that the early differences in the P2 range
are not merely the consequence of latency jitter caused by
stronger variations registered at a later time in each epoch.
Finally, it was shown that the ERP modulations for the fre-
quency and cognate effects were not influenced by repeating
the same stimuli in different blocks.
The time estimate of 172 ms found for the frequency effect is
consistent with the time estimate of 150--175 ms for lexical
access proposed by Indefrey and Levelt (2004). Based on these
authors’ time estimates, both the frequency and the cognate
effect seem to have an early, lexical influence during speech
production. Recall that we used cognate status because it has no
obvious relationship with conceptual variables (see stimulus
ratings) as control for possible conceptual confounds of the
frequency effect. In fact, when plotting cognate and frequency
ERPs together, ERP morphology was remarkably comparable
(see Fig. 6a). Given this similarity and the fact that the cognate
manipulation was not confounded by conceptual variables, we
may interpret the time of split as a good approximation of lexical
access rather than as a difference driven by conceptual factors.
We also need to consider the possibility that the frequency
effect on ERP amplitude may have its origin at a phonological
rather than a lexical level (e.g., Jescheniak and Levelt 1994).
This, however, seems unlikely given that the first significant
ERP differences for both high and low-frequency picture names
and cognate and noncognate picture names occur very early.
Unless one assumes that, once an object is sufficiently
recognized to initiate language related processing (around
150 ms after picture presentation; e.g., Thorpe et al. 1996;
Schmitt et al. 2000), phonological retrieval unfolds in parallel
with lexico-semantic and syntactic retrieval of a word, a pho-
nological account for our findings seems very implausible.
Because the literature does not support the idea that
Figure 4. (a) Low-frequency ERPs compared with high-frequency ERPs inExperiment 1 at PO2 and topographic distribution of electrodes showing a significanteffect at 172 ms after picture presentation (gray area). (b) Noncognate ERPscompared with cognate ERPs in Experiment 1 at PO2 and topographic distribution ofelectrodes showing a significant effect at 200 ms after picture presentation (grayarea).
918 The Time Course of Lexical Access in Speech Production d Strijkers et al.
by guest on Septem
ber 30, 2011cercor.oxfordjournals.org
Dow
nloaded from
phonological and lexical processing proceed in parallel (e.g.,
Salmelin et al. 1994; Van Turennout et al. 1997, 1998; Indefrey
and Levelt 2004), we can conclude that the results here are
indexing lexical access rather than phonological retrieval. Even
more, the present findings do not only show that the first
differences in the ERPs start to emerge at the early P2, but they
also show that the most reliable correlations between ERP
mean amplitude differences and naming latency differences for
the contrasts of interest were present at this component. This
suggests that both these phenomena have an influence during
lexical processing and also that they affect speech production
most strongly at an early point in time, that is, between 160 and
240 ms after picture presentation. In addition, the fact that no
correlations were present between the individual splitting
points in the ERPs for the contrasts at interest and the
individual naming latencies of the fastest conditions of these
contrasts (high-frequency and cognate conditions), suggests
that the time estimate derived from the point of divergence is
indeed indicative for the start (or at least a very early stage) of
lexical access as opposed to its termination. Before considering
the theoretical implications of these findings further, the
robustness and reliability of these results were consolidated by
running a replication experiment. We decided to test Catalan--
Spanish bilinguals performing the same task in their L2
(Spanish) to test whether the timing of the differences would
map onto those found in bilinguals doing the task in their L1. If
the rationale used in the previous experiment is correct we
should be able to replicate these results, regardless of whether
participants name the pictures in their L1 or their L2.
Experiment 2: Catalan--Spanish High Proficient Bilinguals Naming in L2
Method
Participants
Seventeen participants took part in the experiment. All were
highly proficient early Catalan--Spanish bilinguals (a description
of both languages and relative use is presented in Appendix A)
exposed almost exclusively to Catalan during their first 4--5
years of life (see Appendix A) and students at the University of
Barcelona (age range 18--25). Participants received course
credits or monetary compensation for their participation. All
were right-handed, had normal or corrected-to-normal vision
and did not suffer from any neurological or motor problems.
One of the participants had to be removed from the analyses
due to an unacceptably high number of artefacts, leaving 16
participants in the statistical analyses reported below.
Stimuli and Procedures (including EEG acquisition) were
identical to those in Experiment 1.
Analyses
In general, except when specified, all analyses were identical to
those conducted in Experiment 1.
Error Analyses
There were 1.1% erroneous responses, 4.4% outliers, and 3.9%
fast responses which were excluded from the behavioral and
ERP analyses. In addition another 5.3% of the trials were
excluded from the ERP analyses due to artefacts. We decided
not to run a statistical error analysis due to the very low error
rate.
Behavioral Analyses
The behavioral analyses were identical to Experiment 1.
ERP Analyses
The ERP analyses were in all aspects identical to those
described in Experiment 1, except that the time windows
were slightly shifted: 1) 0--80 ms, 2) 80--180 ms, 3) 180--260 ms
(P2), 4) 260--350 ms (N3), and 5) 350--450 ms (P3). In total, ERP
analyses were based on an average of 172 segments (SD = 21)
per condition per subject (low frequency: 173, high frequency:
171, noncognate: 172, cognate: 172).
Figure 5. Spline interpolated grand mean topographies for the differences waves of the frequency (left) and cognate effects (right) at the P2 (split up in 2 time windows of40 ms) in Experiment 1 (upper part; naming in L1) and Experiment 2 (lower part; naming in L2).
Cerebral Cortex April 2010, V 20 N 4 919
by guest on Septem
ber 30, 2011cercor.oxfordjournals.org
Dow
nloaded from
Behavioral Results
Both in the subject and item analyses, significant main effects of
Frequency (F1(1,15) = 45.4, MSE = 2934.7, P < 0.001; F2(1,60) =9.9, MSE = 15014.7, P = 0.003) and Cognate Status (F1(1,15) =33.7, MSE = 2986.2, P < 0.001; F2(1,60) = 7.8, MSE = 15014.7,
P = 0.007) were found. These results replicated those of
Experiment 1 and indicated the presence of frequency and
cognate effects in the naming latencies (see Table 1). The main
effect of Block only reached significance in the item analysis
(F1(5,75) = 2.1, MSE = 10,589.6, P = 0.151; F2(5,300) = 7.3,
MSE = 2397.6, P < 0.001). As in Experiment 1, a significant
interaction between Frequency and Cognate Status was
present for the subject analysis (F1(1,15) = 11.1, MSE = 956.9,
P = 0.005; F2 < 1). The cognate effect was larger for high-
frequency words (43 ms) compared with low-frequency words
(22 ms). Finally, there was a significant interaction between
Frequency and Block in the analysis by subjects (F1(5,75) = 4.9,
MSE = 932.3, P = 0.003; F2(1,60) = 1.6, MSE = 2397.6, P = 0.158)
and a trend toward an interaction between Cognate Status and
Block (F1(5,75) = 2.2, MSE = 851.7, P = 0.081; F2 < 1) only in
the subject analysis. As in Experiment 1 these interactions with
Block did not reveal a stable pattern, but rather randomly
varying sizes of the frequency and cognate effects from block
to block (see Fig. 7). No other interaction reached significance
(P > 0.5).
ERP Results
Onset Latency Analyses
As in Experiment 1, ERPs displayed a typical P1-N1-P2 peak
sequence (see Fig. 8). t-Tests at each sampling rate indicated
that high-frequency ERPs started to diverge significantly from
low-frequency ERPs 184 ms after picture onset (see Figs 8a
and 9a) over anterior, central and posterior midline electrodes
5
Cz-5
Time (ms)
Cz
5
-5
Time (ms)4002000-100 4002000-100
4002000-1004002000-100
Naming in L1 (Experiment 1)
Naming in L2 (Experiment 2)
(b)
(a)
Experiment 1
High FrequencyLow Frequency
Experiment 2
CognateNon-cognate
High Frequency Experiment 1Low Frequency Experiment 1
High Frequency Experiment 2Low Frequency Experiment 2
Cognate Experiment 1Non-Cognate Experiment 1
Cognate Experiment 2Non-Cognate Experiment 2
Ampl
itude
(µV)
Cz
5
-5
Time (ms)
Ampl
itude
(µV)
Cz-5
Time (ms)
5
Cz-5
400Time (ms)
2000-100
5
Figure 6. (a) Low-frequency and high-frequency ERPs compared with noncognate and cognate ERPs at Cz in Experiment 1 (right) and Experiment 2 (left). The frequency ERPsare represented by a full grey and black line. The cognate ERPs are represented by a dotted grey and black line. Negativity is plotted upwards. (b) Between experimentscomparison of the low- and high-frequency ERPs (left), noncognate and cognate ERPs (right), and overall naming in L1 and naming in L2 ERPs (under). Negativity is plottedupwards.
920 The Time Course of Lexical Access in Speech Production d Strijkers et al.
by guest on Septem
ber 30, 2011cercor.oxfordjournals.org
Dow
nloaded from
(at the next time point almost all electrodes showed significant
effects). This time estimate was close to the averaged individual
splitting point (177 ms). The cognate ERPs started to diverge
significantly from noncognate ERPs 184 ms after picture onset
(see Figs 8b and 9b). Furthermore, the time estimate derived
from the grand-averages splitting point overlapped completely
with the averaged individual splitting latencies (184 ms). The
distribution of electrodes showing significant effects at this
time point was widely spread over the scalp (see Fig. 9b). No
significant difference was present between the individual
splitting latency of the frequency effect and that of the cognate
effect (P = 0.4).
Time Window Analyses
Early time windows (0 -- 80 ms and 80 -- 180 ms). None of
the ANOVAs conducted showed significant effects in these
time windows (P > 0.350).
Late time windows (180--260 ms (P2), 260-- 350 ms (N3),
and 350--450 ms (P3). In all 3 time windows significant main
effects were found for Frequency (P2: F(1,15) = 8.9, MSE = 13.6,
P = 0.009; N3: F(1,15) = 74.6, MSE = 23.3, P < 0.001; P3:
F(1,15) = 33.3, MSE = 39.2, P < 0.001) and Cognate Status (P2:
F(1,15) = 10.6, MSE = 12.1, P = 0.005; N3: F(1,15) = 58.3, MSE =22.3, P < 0.001; P3: F(1,15) = 20.6, MSE = 28.5, P < 0.001). ERPs
in the high-frequency condition were significantly more
negative going compared with those elicited by the low-
frequency condition and ERPs in the cognate condition were
significantly more negative going than those in the noncognate
condition. Both effects were widely distributed over the scalp,
reaching their maximum at posterior and right-frontal sites
(see Fig. 5). The interaction between Frequency and Cognate
Status was significant, but only for the 2 later time windows
(P2: F < 1; N3: F(1,15) = 18.8, MSE = 14.1, P = 0.001; P3:
F(1,15) = 20.9, MSE = 11.5, P < 0.001). It is noteworthy that this
pattern of results is identical to that found in Experiment 1.
Finally, as in Experiment 1, in none of the 3 time windows
there was a significant effect of Block (P > 0.250), or
interactions between Block and Frequency (P > 0.150) or
Cognate Status (F < 1).
Correlation Analyses
As in Experiment 1, no significant correlations were found
between the individual splitting point of the frequency and
cognate contrasts in the ERPs with the individual mean naming
latencies of those contrasts in the behavioral data (P > 0.4), nor
did we find significant correlations between the individual peak
latencies of the P2, N3, and P3 (P > 0.36).
Significant positive Pearson product--moment correlations
were found between P2 mean amplitude difference between
high and low frequency and difference in naming latencies
between the high- and low-frequency condition at fronto-
central (R = 0.512, P = 0.042) and left central electrodes (R =0.492, P = 0.053). There was also a trend towards a positive
correlation at centro-parietal electrodes (R = 0.447, P = 0.082).
In the N3 range marginally significant positive correlations
between the frequency effect in the naming latencies and the
frequency effect in the ERPs were present at left parietal (R =0.492, P = 0.053) and left central electrodes (R = 0.483, P =0.058). Small trends toward positive correlations in the same
direction were present at centro-parietal (R = 0.465, P = 0.070)
and right central electrode clusters (R = 0.427, P = 0.099).
Finally, for the P3, only a small trend towards a positive
correlation was present at left parietal electrode sites (R =0.428, P = 0.098).
For the cognate contrast a similar pattern of correlations was
found: There were significant correlations between the
difference in P2 mean amplitude between cognate and
noncognate conditions and the difference in naming latencies
between cognate and noncognate conditions at left frontal (R =0.535, P = 0.033) and left central electrode clusters (R = 0.502,
P = 0.047). A trend towards a positive correlation was present
at fronto-central electrodes (R = 0.449, P = 0.081). As in
Experiment 1 these correlations disappeared in the N3 (P >
0.190) and the P3 (P > 0.560) range, respectively.
Comparison with Experiment 1
Differences in ERP splitting point latencies and RTs between
experiments (i.e., between the 2 participant groups) were not
significant (all P > 0.1). However, we did observe a marginally
significant main effect in the P2 range (and also for subsequent
peaks) between Groups (naming in L1 vs. naming in L2;
F(1,30) = 3.4, MSE = 77.1, P = 0.060; see Fig. 6b) and,
importantly, a marginal significant interaction between Group
and Frequency (F(1,30) = 3.8, MSE = 7.2, P = 0.062; see Fig. 6b).
The frequency effect showed a larger amplitude difference at
the P2 in Experiment 2 (naming in L2; difference: 1.9 lV)compared with Experiment 1 (naming in L1; difference: 1.1 lV).None of the other interactions with Group turned out
significant (all P > 0.1).
Discussion
The results of Experiment 2 in a different group of participants
performing the picture naming task in their L2 were overall
highly similar to those obtained in Experiment 1. First, naming
latencies displayed the expected frequency and cognate
effects, and the interaction between frequency and cognate
status was also replicated. (At the moment we do not have an
explanation for this interaction. A similar interaction has been
reported before in behavioural picture naming experiments,
e.g., Ivanova and Costa 2008, but also the reverse interaction;
e.g., Christoffels et al. 2003, and sometimes none; e.g., Costa
et al. 2000. It might be that the presence and direction of this
620
660
700
740
780
820
1 2 3 4 5 6
Blocks
Nam
ing
Late
ncie
s (m
s)
LFNC
LFC
HFNC
HFC
Figure 7. Mean naming latencies in Experiment 2 (LFNC 5 low-frequencynoncognate; LFC 5 low-frequency cognate; HFNC 5 high-frequency noncognate;HFC 5 high-frequency cognate) over repetitions.
Cerebral Cortex April 2010, V 20 N 4 921
by guest on Septem
ber 30, 2011cercor.oxfordjournals.org
Dow
nloaded from
interaction depends on differences of [uncontrolled] stimulus
properties between experiments. Independently, in both
experiments the interaction becomes apparent in the ERPs
only after the splitting point and the P2 [i.e., after 240 ms],
which is beyond the period of interest in this paper.) Second, in
the ERPs, early differences between low and high frequency
ERPs (184 ms) and between cognate and noncognate ERPs
(184 ms) were observed in a similar time window as those
found in experiment 1. Third, the pattern of correlations
resembled that seen in Experiment 1. Indeed, there were no
correlations between the individual splitting points of the
experimental contrasts and the individual naming latencies of
the fastest condition of those contrasts. There were no
correlations between P2, N3, and P3 peak latencies either,
but significant positive correlations between individual fre-
quency and cognate effects in the naming latencies with mean
amplitude differences for both the frequency and cognate
effects in the ERPs. These correlations were strongest in the P2
range and weaker or absent in the N3 and P3 ranges.
Only 2 qualitative differences were found between the 2
experiments:
1) The frequency effect had an earlier onset compared with
the cognate effect in Experiment 1, but this latency difference
was absent in Experiment 2. Faster conceptual processing for
high frequency words due to their higher familiarity (a
conceptual property), which was not present for the cognate
F3 Fz F4
FC1 FC2
C3 Cz C4
CP1 CP2
P3 Pz P4
PO1 PO2
O1 Oz O2
-5 [µV]
P2
P3
N3
High Frequency Low Frequency
F3 Fz F4
FC1 FC2
C3 Cz
CP1 CP2
P3 Pz
PO1 PO2
O1 Oz O2
Cognates Non-Cognates
(a)
(b)
C4
P4
-5 [µV]
200 550400
200 550400
Figure 8. (a) Low-frequency ERPs compared with high-frequency ERPs in Experiment 2 at anterior, central, and posterior scalp locations. Low-frequency ERPs are representedby a dotted line and high frequency ERPs by a full line. Negativity is plotted upwards. (b) Noncognate ERPs compared with cognate ERPs in Experiment 2 at anterior, central, andposterior scalp locations. Noncognate ERPs are represented by a dotted line and Cognate ERPs by a full line. Negativity is plotted upwards.
922 The Time Course of Lexical Access in Speech Production d Strijkers et al.
by guest on Septem
ber 30, 2011cercor.oxfordjournals.org
Dow
nloaded from
contrast, could account for the faster engagement in lexical
processing for those items and consequently for the earlier
splitting latency. However, the similarity in splitting latency for
lexical frequency and cognate status in Experiment 2 invalidates
this account. In addition, given that analyses of splitting latencies
for lexical frequency and cognate status are based on different
sets of stimuli (50% different), it is likely that conceptual
processing has a different duration on average for each picture
set (in that sense the similarity in Experiment 2 is more sur-
prising than the different onset in Experiment 1). Nevertheless,
ERPs had the same morphology in both conditions (see Fig. 6a)
and amplitude differences were found in the time window
previously associated with lexical processing (175--250 ms
according to Indefrey and Levelt 2004).
Second, we encountered more positive P2 amplitudes in
Experiment 2 (naming in L2) as compared with Experiment 1
(naming in L1; see Fig. 6b). This is an important observation in
light of our theoretical claims. (This observation is also of
importance regarding the bilingual naming disadvantage in the
non dominant language; e.g., Indefrey 2006; Ivanova and Costa
2008). However, because this topic does not fall under the
scope of the present study, this result will only be discussed in
light of the theoretical claims made in the present study. These
findings with respect to the bilingual naming disadvantage will
be discussed elsewhere [including more subjects in each group
and adding a within-subjects experiment].) One may argue that
due to the interactivity of the brain, different representational
systems which are interconnected, such as the lexical system
and the semantic (or object—imaginal—representation)
system, may benefit and even share processing activation from
one domain to the other (e.g., Paivio 1986). In such a scenario
our results would still reflect the first influence of lexical
variables during speech production processing (given the
cognates), but not necessarily processing solely at a lexical
level. This being said, the dual-code view cannot, however,
account for the amplitude difference between L1 and L2
naming in the P2 range (see Fig. 6b). Naming in L1 versus
L2 should activate the exact same semantic (object) represen-
tation (e.g., Kroll and Stewart 1994) and a difference between
the 2 can only be explained at the lexical level where the
representational format is distinct (recall that subjects are
early, highly proficient bilinguals using both languages on
a daily basis, and that stimuli were concrete words). Although
the validity of between group comparisons can be disputed
(especially with ERPs), it is difficult to imagine that 2 groups of
participants viewing the exact same images overall would
display between-group differences by chance in the same time
range (~192 ms) and in the same manner as differences
generated by lexical frequency and cognate status manipu-
lations. This is especially true because we also encountered
a (marginally) significant interaction with lexical frequency
between experiments. Such pattern of results is unlikely to
sprout from coincidental between-group differences. A stron-
ger P2 modulation for the frequency effect during L2 naming
compared with L1 naming can only be readily explained by
assuming that these effects occur at the lexical level. In
addition, the fact that for lexical frequency there was
a difference in familiarity (a conceptual property) while this
difference was absent for cognate status, and that both
variables elicited similar ERP effects, also argues against the
possibility that present results merely reflect lexical influences
during conceptual processing.
General Discussion
The main aim of this study was to characterize the time course
of lexical access in speech production using a high-resolution
temporal technique, event-related potentials. Two effects
known to affect picture naming latencies were investigated:
the word frequency and the cognate effects. Participants
showed reliable and robust frequency and cognate effects in
both experiments, replicating previous studies (e.g., Oldfield
and Wingfield 1965; Jescheniak and Levelt 1994; Costa et al.
2000; Navarrete et al. 2006; Almeida et al. 2007; Christoffels
et al. 2007). Crucially, we found early ERP effects of frequency
and cognate status, independently of whether naming was
Figure 9. (a) Low-frequency ERPs compared with high-frequency ERPs inExperiment 2 at PO2 and topographic distribution of electrodes showing a significanteffect at 184 ms after picture presentation (grey area). (b) Noncognate ERPscompared with cognate ERPs in Experiment 1 at PO2 and topographic distribution ofelectrodes showing a significant effect at 184 ms after picture presentation (greyarea).
Cerebral Cortex April 2010, V 20 N 4 923
by guest on Septem
ber 30, 2011cercor.oxfordjournals.org
Dow
nloaded from
performed in L1 or L2. High-frequency ERPs diverged from
low-frequency ERPs at around 180 ms after picture onset
(172 ms in Experiment 1, 184 ms in Experiment 2) coinciding
with the onset of a positive wave (P2), with the high lexical
frequency condition eliciting lower ERP amplitudes than the
low-frequency condition. A similar pattern of results was found
for cognates: pictures with cognate names started to diverge
from those elicited by pictures with noncognate names around
190 ms after stimulus presentation (200 ms in Experiment 1;
184 ms in Experiment 2), with noncognates eliciting greater
amplitudes than cognates.
The Word Frequency Effect as an Index of Lexical Access
The early effect of word frequency in picture naming suggests
that speakers start the lexicalization process very early on after
picture presentation. That is, to the extent that word frequency
affects lexical retrieval, we propose that participants started the
lexicalization processes between 150 and 200 ms after picture
onset. This time estimation is consistent with results from MEG
studies (e.g., Levelt et al. 1998; Maess et al. 2002) and covert
‘‘lexical’’ ERP studies (e.g., Schmitt et al. 2000), and fundamen-
tally agrees with the meta-analysis conducted by Indefrey and
Levelt (2004). Importantly, the early effect of word frequency
during picture naming is unlikely to index the time of retrieval
of target lexical representations given the absence of correlation
between ERP splitting latencies and naming latencies for high-
frequency words. Instead, we propose that this effect coincides
with initial activation and retrieval operations within the lexicon.
That is, the point in time where a lexical representation gets
activated and enters in the competitive process of selection,
with high-frequency items showing a head start over low-
frequency items due to their (permanently) higher activation
levels. The splitting point between ERPs can therefore be seen as
the transition phase between conceptual and lexical selection
processes (see e.g., Thorpe et al. 1996; Hauk et al. 2007; for
object recognition time estimates).
Besides the descriptive chronometric information provided
by our results, the present findings also have implications for
the locus of the frequency effect in speech production. As
mentioned in the introduction, word frequency affects solely
the retrieval of the phonological properties of a word accord-
ing to some authors (e.g., Jescheniak and Levelt 1994;
Jescheniak et al. 2003). Under this assumption, we should
have expected the ERPs in the high- and low-frequency
conditions to start diverging when a word’s phonological code
is supposed to be retrieved, that is, around 275 ms according to
Indefrey and Levelt (2004). The presence of a word frequency
effect at ~180 ms is at odds with this position, and suggests that
word frequency also affects the speed with which lexical items
are retrieved from the lexicon (e.g., Caramazza et al. 2001;
Navarrete et al. 2006; Almeida et al. 2007).
However, finding an early effect of word frequency does not
discard an effect at later processing stages, such as that of
phonological encoding. Indeed, our results show that differ-
ences between low-frequency and high-frequency ERPs are
present in later time windows as well. In fact, the amplitude of
the ERPs for high- and low-frequency words correlated
positively with naming latency differences for time windows
at which lexical (the P2 window) but also phonological
encoding (the N3 window) are supposed to take place.
Therefore, considering hypothetically that different cognitive
processes take place in these different time windows (e.g.,
lexical access -early stage-, phonological access -late stage-), we
can conclude that both processing stages are affected by word
frequency. This view of the ubiquitous effects of word
frequency is consistent with recent hemodynamic evidence
showing that word frequency modulates the activity of brain
areas thought to be involved in the retrieval of lexico-semantic
as well as phonological information (Graves et al. 2007) and
recent studies with brain-damaged patients showing frequency
effects for semantically and phonologically related errors and
errors resulting in nonwords (e.g., Kittridge et al. 2007; Knobel
et al. 2008).
A possible caveat when interpreting the early effect of word
frequency in the ERP data is the potential correlation of this
variable with conceptual variables such as familiarity and
imageability. Indeed, stimuli ratings on conceptual variables
conducted for the pictures used in the present experiments
showed significant familiarity differences between the low-
frequency and high-frequency items. This means that the early
ERP effects for frequency could be driven by these conceptual
differences. However, we also found early cognate effects in
the ERPs, in a rather similar time window to that of the
frequency effects. In fact, when plotted together the electro-
physiological signature of the cognate effect and the frequency
effect are practically identical (see Fig. 6a) and the correlation
patterns between behavioral differences and electrophysiolog-
ical differences are also similar. Given that -unlike word
frequency cognate status is not correlated with conceptual
variables (see also stimuli ratings), the early effect of cognate
status cannot be interpreted as a mere effect of correlated
conceptual variables, but rather some sort of lexical effect.
Thus, if one is willing to interpret the cognate effect at such
early time as revealing lexical processing, it is reasonable to
interpret word frequency effects along the same lines.
On the Origin of the Cognate Effect
As mentioned above, the results of the cognate manipulation are
useful when interpreting the word frequency effects. In
addition, the early effect of cognate status also sheds light on
its origin. Interestingly, such an effect was also descriptively
reported in the study by Christoffels et al. (2007, personal
communication), who found significant differences between
cognate and noncognate ERPs as early as 175 ms after picture
presentation.
The parallel results observed for word frequency and
cognate manipulations suggest that the 2 effects might have
the same origin at the lexical level. Because of the phonological
overlap between a cognate and its translation, every time
a cognate word is heard or uttered, both the target lexical
representation and its translation are strongly activated,
irrespective of the language of utterance. In contrast, when
a noncognate word is produced or heard, the translation word
will probably not be activated that strongly, given the lack of
phonological overlap. Following this rationale, cognate lexical
representations should have a higher frequency than non-
cognate lexical representations, because the former are
activated more often. (Both interactive, e.g., Dell 1986, as
sequential, e.g., Levelt, et al. 1998, models of speech production
can nicely capture this assumption. According to interactive
models, the activated phonological segments of the target word
will send activation back to any word with which they are
924 The Time Course of Lexical Access in Speech Production d Strijkers et al.
by guest on Septem
ber 30, 2011cercor.oxfordjournals.org
Dow
nloaded from
linked. In such scenario, the utterance of a cognate word will
cause its translation to become lexical activated as well, due to
feedback activation from shared phonological segments onto
the lexical level with which these segments are linked. A
similar process will not unfold for noncognate translations,
because they hardly share phonological segments; see Costa
et al. 2005. Sequential models can explain a cognate effect at
the lexical level in a similar manner through comprehension:
every time a cognate is heard [also through perception of one’s
own voice], the similar phonological content will also activate
the nontarget lexical representation, whereas for noncognates
this bottom-up activation via phonology will only result in
activation of the target word.) Consequently, the cognate effect
may reflect a word frequency effect in disguise, with cognate
words behaving as high-frequency words and noncognate
words as low-frequency words (see also Kirsner et al. 1993;
Sanchez-Casas and Garcıa-Albea 2005, for an alternative
explanation of cognate effects at the lexical level). Note that
a similar explanation at the conceptual level cannot account for
the cognate effect because the conceptual representations of
cognates and noncognates are considered to be shared
between L1 and L2 (e.g., Kroll and Stewart 1994).
Independently of the precise nature of the cognate effect,
our results reveal that cognate status has an effect at early
stages of lexical access. However, as in the case of word
frequency, this early effect does not preclude effects of
cognate status at subsequent processing levels. For instance,
Christoffels et al. (2007) reported ERP cognate effects between
275 and 375 ms after target presentation, with enhanced
negative amplitude for cognate relative to noncognate ERPs.
Consistent with this late effect, we found a similar modulation
at the N3. Thus, as reported previously for word frequency,
cognate status appears to affect both lexical processing (P2
range) and phonological encoding (N3 range), and therefore
seems to affect picture naming latencies in a similar way as
word frequency.
Methodological Issues and ERP Components of Interest
So far we have mainly discussed the data in the temporal
domain. Given the novelty of EEG studies using overt picture
naming, it is pertinent to dedicate some words to the ERP
components of interest identified in the experiments. Before
doing so, however, some potential methodological pitfalls need
to be discussed.
In the experiments there were 16 stimuli per condition (32
per experimental contrast), and many repetitions were
needed (6) in order to obtain enough trials per condition.
Stimulus repetition in ERPs can be a substantial source of
modulation (e.g., Bentin and McCarthy 1994; Rugg and Doyle
1994). The repetition effects in this study were however
negligible and, critically, there were no interactions between
repetition and other factors in either experiments, suggesting
that stimulus repetition did not affect the frequency and
cognate effects reported. In particular, the P2 modulation by
frequency and cognate status had the same magnitude in all
experimental blocks. The small magnitude of repetition
effects observed, may be due to 1) the fact that we did not
record the first presentation of the stimuli (familiarization
phase). Therefore we could not measure the ERP differences
between the familiarization phase and the first experimental
presentation, where the strongest ERP repetition effects are
to be expected. Indeed, ERP studies using multiple repetitions
have shown that ERP differences elicited by repetition are
largest for the first repetition and seem to decrease or even
vanish with subsequent repetitions (e.g., Gruber and Muller
2005; Friedman and Cycowicz 2006); 2) the irregularity of the
lag between repeated items (e.g., Henson et al. 2004); 3) the
relatively large average lag between repeated stimuli (e.g.,
Henson et al. 2004).
Another possible confound in the present study is that
different physical stimuli were used in the different experi-
mental conditions. This could result in spurious ERP amplitude
modulations caused by physical stimulus differences rather
than the cognitive manipulation of interest (e.g., Picton et al.
2000). However, because 50% of the stimuli were completely
different between the high-frequency and cognate condition,
finding similar time courses of differences and correlation
patterns by chance is unlikely. In addition, the ERP pattern
correlated with the behavioral results, where naming latency
differences are less likely to sprout from distinct physical
stimuli. Finally, when comparing directly L1 against L2 naming
overall, that is, when comparing ERPs elicited by the exact
same set of pictures in 2 different naming contexts, we find the
same P2 modulation as in experiments 1 and 2 taken separately
(see Fig. 6b).
The ERP components observed here were not systematically
interpreted in a traditional way because this study focused on
divergence latencies and amplitude-naming latency correlations.
Nevertheless, the peaks observed may be related to classical
components described in the literature. For instance, P3 mean
amplitude was more pronounced for low- than for high-
frequency words, which may suggest postlexical reprocessing
of word-related information (e.g., Polich and Donchin 1988;
Hauk and Pulvermuller 2004). However, given the marked
positivity of the P3 component in our dataset, amplitude
modulation could also partially reflect early stages of motor
preparation.
Perhaps the most interesting results were found for the P2
range, given the correlation between frequency and cognate
naming latency and P2 mean amplitude difference between
high- and low-frequency picture names and between cognates
and noncognates. Individuals showing larger frequency and
cognate naming latency effects also showed bigger P2 mean
amplitude differences for the same contrasts. In other words, P2
amplitude appears to reflect the ease of lexical access, with
lower amplitudes associated with easily accessible representa-
tions (high frequency words and cognates) and larger ampli-
tudes associated with less accessible representations. Such
amplitude modulations are consistent with Hebbian theory of
cell assemblies (e.g., Pulvermuller 1999; Hauk and Pulvermuller
2004). Another possible interpretation for the P2 comes from
word recognition experiments manipulating vocabulary class
(e.g., King and Kutas 1998; Brown et al. 1999; Osterhout et al.
2002). These studies reported, in a similar vein as observed in
the present study, reduced P2 amplitudes for closed class (faster
condition) as compared with open class words (slower
condition). Although these studies did not elaborate much on
this finding, it was suggested that the P2 modulation might
reflect attentional differences between nonlexical aspects of the
stimuli such as length (cf., Mangun and Hillyard 1995). For the
present study however, such an account does not seem valid.
Given the results observed for the cognates, as explained
extensively above, the P2 modulations reported here have to be
Cerebral Cortex April 2010, V 20 N 4 925
by guest on Septem
ber 30, 2011cercor.oxfordjournals.org
Dow
nloaded from
related, at least in part, to linguistic processes. It is possible that
these P2 effects are indeed confounded by attention, with rare
stimuli eliciting larger attentional shifts than more common
stimuli (e.g., Luck and Hillyard 1994), but this would not take
away the value of our observation because, in that case, these P2
differences most likely reflect attentional resources needed
during lexical activation. Future research exploring further the
functional characteristics of the P2 component will probably
provide fundamentally new insights regarding lexical access in
speech production.
Conclusion
For the first time, early electrophysiological differences elicited
by manipulation of lexical frequency and cognate status during
overt picture naming were established. Based on the latency of
ERP divergences between conditions, lexical access is esti-
mated to occur at around 180 ms after target presentation.
Aside from this important chronometric contribution, the
present study offers a promising new paradigm using a simple
task and a simple experimental design to study the time course
of speech production. In addition, by identifying an early
electrophysiological correlate of lexical processing, this study
may be a starting point for approaching a variety of
psycholinguistic phenomena in language production from
a whole new perspective.
Funding
Spanish Government grant (PSI2008-01191); Project Consolider-
Ingenio 2010 (CSD 2007-00012); and Spanish Government
(FPU-2007-2011) predoctoral fellowship supported K.S.
Notes
We would like to thank Phillip Holcomb and 2 anonymous reviewers for
their helpful comments to the previous version of this manuscript. We
would also like to thank Elin Runnqvist for her help in revising this
manuscript. Conflict of Interest : None declared.
Address correspondence to Albert Costa, PhD, Dept. de Tecnologia,
ICREA, Universitat de Pompeu Fabra, C/ Tanger, 122-140, 08018
Barcelona, Spain. Email: [email protected].
Appendix A. Language history and the self-assessed proficiency forall participants
Language history and self-assessed proficiency scores of participants.
Mean age and SD are given in years. The onset of L2 acquisition refers
to the mean age (in years) at which participants started learning
Catalan/Spanish. ‘‘Use of L2’’ refers to how long (in years) participants
have been using the L2 regularly. The proficiency scores were obtained
through a questionnaire filled out by the participants after the
experiment. The scores are on a 4 point scale, in which 4 represents
native speaker level; 3, good level; 2, medium level; and 1, poor level of
proficiency. The self-assessed index is the average of the participants’
responses to 4 domains (speech comprehension, speech production,
reading, and writing).
Appendix B: List of stimuli used in the Experiment
References
Almeida J, Knobel M, Finkbeiner M, Caramazza A. 2007. The locus of the
frequency effect in picture naming: when recognizing is not
enough. Psychon Bull Rev. 14(6):177--1182.
Bentin S, McCarthy G. 1994. The effect of immediate stimulus
repetition on reaction time and event-related potentials in tasks of
different complexity. J Exp Psychol Learn Mem Cogn. 20:130--149.
Brown CM, Hagoort P, ter Keurs M. 1999. Electrophysiological
signatures of visual lexical processing: open and closed-class words.
J Cogn Neurosci. 11:261--281.
Caramazza A. 1997. How many levels of processing are there in lexical
access? Cognit Neuropsychol. 14:177--208.
Caramazza A, Costa A, Miozzo M, Bi Y. 2001. The specific-word
frequency effect: implications for the representation of homo-
phones. J Exp Psychol Learn Mem Cogn. 27:1430--1450.
Christoffels IK, Firk C, Schiller NO. 2007. Bilingual language control: an
event-related brain potential study. Brain Res. 1147:192--208.
Coles MGH. 1989. Modern mind-brain reading: psychophysiology,
physiology and cognition. Psychophysiology. 26:251--269.
Costa A, Caramazza A, Sebastian-Galles N. 2000. The cognate facilitation
effect: implications for models of lexical access. J Exp Psychol Learn
Mem Cogn. 26:1283--1296.
Costa A, Santesteban M, Cano A. 2005. On the facilitatory effects of cog-
nate words in bilingual speech production. Brain Lang. 94:94--103.
Christoffels IK, De Groot AMB, Waldorp LJ. 2003. Basic skills in
a complex task: a graphical model relating memory and lexical
retrieval to simultaneous interpreting. Biling Lang Cogn. 6:201--211.
Dell GS. 1986. A spreading-activation theory of retrieval in sentence
production. Psychol Rev. 93:283--321.
Dell GS. 1990. Effects of frequency and vocabulary type on phonolog-
ical speech errors. Lang Cogn Proc. 5:313--349.
Language history Age L2 onset L2 use # Years in Catalunya
Spanish--Catalan bilinguals 20 (2) 4 (2) 16 (4) 20 (1)Catalan--Spanish bilinguals 22 (2) 5 (3) 17 (2) 22 (1)
L1 L2Self-assessed proficiencySpanish--Catalan bilinguals 3.92 3.51Catalan--Spanish bilinguals 3.94 3.68
Low-frequency noncognates Low-frequency cognates
Spanish Catalan English Spanish Catalan EnglishLagartija Sargantana [Lizard] Cocodrilo Cocodril [Alligator]Melocoton Pressec [Peach] Platano Platan [Banana]Hucha Guardiola [Piggybank] Escoba Escombra [Broom]Peonza Baldufa [Top] Guante Guant [Glove]Calcetın Mitjo [Sock] Dragon Drac [Dragon]Muela Queixal [Tooth] Elefante Elefant [Elephant]Zanahoria Pastanaga [Carrot] Martillo Martell [Hammer]Tenedor Forquilla [Fork] Raton Ratolı [Mouse]Buho Mussol [Owl] Pincel Pinzell [Paintbrush]Pato Anec [Duck] Pinguino Pinguı [Penguin]Cepillo Raspall [Brush] Patın Patı [Roller skate]Cubo Galleda [Bucket] Cuchara Cullera [Spoon]Hacha Destral [Axe] Tanque Tanc [Tank]Rana Granota [Frog] Violın Violı [Violin]Grifo Aixeta [Faucet] Trineo Trineu [Sled]Ardilla Esquirol [Squirrel] Tigre Tigre [Tiger]
High-frequency noncognates High-frequency cognates
Spanish Catalan English Spanish Catalan EnglishManzana Poma [Apple] Nube Nuvol [Cloud]Queso Formatge [Cheese] Oreja Orella [Ear]Naranja Taronja [Orange] Plato Plat [Plate]Cerdo Porc [Pig] Flor Flor [Flower]Cuchillo Ganivet [Knife] Arbol Arbre [Tree]Rama Branca [Branch] Gato Gat [Cat]Huevo Ou [Egg] Caja Caixa [Box]Hoja Fulla [Leaf] Banco Banc [Bench]Sombrero Barret [Hat] Avion Avio [Airplane]Bolsillo Butxaca [Pocket] Reloj Rellotge [Watch]Silla Cadira [Chair] Nariz Nas [Nose]Lluvia Pluja [Rain] Caballo Cavall [Horse]Perro Gos [Dog] Brazo Bracx [Arm]Ojo Ull [Eye] Telefono Telefon [Telephone]Ventana Finestra [Window] Pie Peu [Foot]Mesa Taula [Table] Libro Llibre [Book]
926 The Time Course of Lexical Access in Speech Production d Strijkers et al.
by guest on Septem
ber 30, 2011cercor.oxfordjournals.org
Dow
nloaded from
Dell GS, O’Seaghdha PG. 1991. Mediated and convergent lexical priming
in language production: a comment on Levelt et al. Psychol Rev.
98:604--614.
Eddy M, Schmid A, Holcomb PJ. 2006. Masked repetition priming and
event-related brain potentials: a new approach for tracking the time
course of object perception. Psychophysiology. 43:564--568.
Eimer M. 1993. Effects of attention and stimulus probability on ERPs in
a go/no-go task. Biol Psychol. 35:123--138.
Eulitz C, Hauk O, Cohen R. 2000. Electroencephalographic activity over
temporal brain areas during phonological encoding in picture
naming. Clin Neurophysiol. 111:2088--2097.
Friedman D, Cycowicz YM. 2006. Repetition priming of possible and
impossible objects from ERP and behavioural perspectives. Psycho-
physiology. 43:569--578.
Ganushchak LY, Schiller NO. 2008. Motivation and semantic context
affect brain error-monitoring activity: an event-related brain
potentials study. Neuroimage. 39:395--405.
Grainger J, Holcomb PJ. 2009. Watching the word go by: on the time
course of component processes in visual word recognition. Lang
Linguist Compass. 3:128--156.
Gratton G, Coles MGH. 1989. Generalization and evaluation of eye-
movement correction procedures. J Psychophysiol. 3:14--16.
Graves WW, Grabowski TJ, Mehta S, Gordon JK. 2007. A neural
signature of phonological access: distinguishing the effects of word
frequency from familiarity and length in overt picture naming. J
Cogn Neurosci. 19:617--631.
Gruber T, Muller MM. 2005. Oscillatory brain activity dissociates
between associative stimulus content in a repetition priming task in
the human EEG. Cereb Cortex. 15:109--116.
Guthrie D, Buchwald JS. 1991. Significance testing of difference
potentials. Psychophysiology. 28:240--244.
Hauk O, Patterson K, Woollams A, Cooper-Pey E, Pulvermuller F,
Rogers TT. 2007. How the camel lost its hump: the impact of object
typicality on event-related potential signals in object decision. J
Cogn Neurosci. 19:1338--1353.
Hauk O, Patterson K, Woollams A, Watling L, Pulvermuller F, Rogers TT.
2006. [Q:] When would you prefer a SOSSAGE to a SAUSAGE? [A:] At
about 100 msec ERP correlates of orthographic typicality and
lexicality in written word recognition. J Cogn Neurosci. 18:818--832.
Hauk O, Pulvermuller F. 2004. Effects of word length and frequency on
the human event-related potential. Clin Neurophysiol.
115:1090--1103.
Hauk O, Pulvermuller F, Ford M, Marslen-Wilson WD, Davis MH. 2009.
Can I have a quick word? Early electrophysiological manifestations
of psycholinguistic processes revealed by event-related regression
analysis of the EEG. Biol Psychol. 80:64--74.
Henson RNA, Rylands A, Ross E, Vuilleumeir P, Rugg MD. 2004. The
effect of repetition lag on electrophysiological and haemodynamic
correlates of visual object priming. Neuroimage. 21:1674--1689.
Holcomb PJ, Grainger J. 2006. The time course of masked repetition
priming: an event-related brain potential investigation. J Cogn
Neurosci. 18:1631--43.
Holcomb PJ, McPherson WB. 1994. Event-related brain potentials
reflect semantic priming in an object decision task. Brain Cogn.
24:259--276.
Ivanova I, Costa A. 2008. Does bilingualism hamper lexical access in
speech production? Acta Psychol. 127:277--288.
Indefrey P. 2006. A meta-analysis of hemodynamic studies on first and
second language processing: which suggested differences can we
trust and what do they mean? Lang Learn. 56:279--304.
Indefrey P, Levelt WJM. 2004. The spatial and temporal signatures of
word production components. Cognition. 92:101--144.
Jescheniak JD, Levelt WJM. 1994. Word frequency effects in speech
production: retrieval of syntactic information and of phonological
form. J Exp Psychol Learn Mem Cogn. 20:824--843.
Jescheniak JD, Meyer AS, Levelt WJM. 2003. Specific-word frequency is
not all that counts in speech production: comments on Caramazza,
Costa et al. (2001) and new experimental data. J Exp Psychol Learn
Mem Cogn. 29:432--438.
Jescheniak JD, Schriefers H, Garrett MF, Friederichi AD. 2002. Exploring
the activation of semantic and phonological codes during
speech planning with event-related brain potentials. Brain Lang.
94:94--103.
Johnson JS, Olshausen BA. 2005. The earliest EEG signatures of object
recognition in a cued-target task are postsensory. J Vis. 5:299--312.
Kiefer M. 2001. Perceptual and semantic sources of category-specific
effects: event-related potentials during picture and word categori-
zation. Mem Cognit. 29:100--116.
King JW, Kutas M. 1998. Neural plasticity in the dynamics of human
visual word recognition. Neurosci Lett. 2:244--261.
Kirsner K, Lalor E, Hird K. 1993. The bilingual lexicon: exercise,
meaning and morphology. In: Schreuder R, Weltens B, editors. The
bilingual lexicon. Amsterdam: John Benjamins. p. 215--246.
Kittridge AK, Dell GS, Verkuilen J, Schwartz MF. 2007. Where is the
effect of frequency in word production? Insights from aphasic
picture-naming errors. Cognit Neuropsychol. 1:1--30.
Koester D, Schiller N. 2008. Morphological priming in overt language
production: electrophysiological evidence from Dutch. Neuro-
image. 42:1622--1630.
Kok A. 1986. Effects of degradation of visual stimuli on components of
the event-related potential (ERP) in go/nogo reaction tasks. Biol
Psychol. 23:21--38.
Knobel M, Finkbeiner M, Caramazza A. 2008. The many places of
frequency: evidence for a novel locus of the lexical frequency effect
in word production. Cognit Neuropsychol. 25:256--286.
Kroll JF, Stewart E. 1994. Category interference in translation and
picture naming: evidence for asymmetric connections between
bilingual memory representations. J Mem Lang. 33:149--174.
Levelt WJM, Praamstra P, Meyer AS, Helenius P, Salmelin R. 1998. A MEG
study of picture naming. J Cogn Neurosci. 10:553--567.
Levelt WJM, Roelofs A, Meyer AS. 1999. A theory of lexical access in
speech production. Behav Brain Sci. 22:1--75.
Luck SJ, Hillyard SA. 1994. Electrophysiological correlates of feature
analysis during visual search. Psychophysiology. 31:291--308.
Maess B, Friederici AD, Damian M, Meyer AS, Levelt WJM. 2002.
Semantic category interference in overt picture naming: an MEG
study. J Cogn Neurosci. 14:455--462.
Mangun GR, Hillyard SA. 1995. Mechanisms and models of selective
attention. In: Rugg MD, Coles MGH, editors. Electrophysiology of
mind. Oxford: Oxford University Press. p. 40--85.
Masaki H, Tanaka H, Takasawa N, Yamazaki K. 2001. Error-related brain
potentials elicited by vocal errors. Cogn Neurosci Neurorep.
12:1851--1855.
Miller J, Riehle A, Requin J. 1992. Effects of preliminary perceptual
output on neuronal activity of the primary motor cortex. J Exp
Psychol Hum Percept Perform. 18:1121--1138.
Navarrete E, Basagni B, Alario XF, Costa A. 2006. Does word frequency
affect lexical selection in speech production? Q J Exp Psychol.
10:1681--1690.
Oldfield RC, Wingfield A. 1965. Response latencies in naming objects. Q
J Exp Psychol. 17:273--281.
Osterhout L, Allen M, McLaughlin J. 2002. Words in the brain: lexical
determinants of word-induced brain activity. J Neurolinguist.
15:171--187.
Pallier C, Dupoux E, Jeannin X. 1997. EXPE: An expandable pro-
gramming language for on-line psychological experiments. Behav
Res Methods. 29:322--327.
Pfefferbaum A, Ford JM, Weller BJ, Kopell BS. 1985. ERPs to response
production and inhibition. Clin Neurophysiol. 60:423--434.
Picton TW, Bentin S, Berg P, Donchin E, Hillyard SA, Johnson R, Jr,
Miller GA, Ritter W, Ruchkin DS, Rugg MD, et al. 2000. Guidelines
for using human event-related potentials to study cognition:
recording standards and publication criteria. Psychophysiology.
37:127--152.
Polich J, Donchin E. 1988. P300 and the word frequency effect.
Electroencephalogr Clin Neurophysiol. 70:33--45.
Pulvermuller F. 1999. Words in the brain’s language. Behav Brain Sci.
22:253--79.
Rodriguez-Fornells A, Schmitt BM, Kutas M, Munte TF. 2002.
Electrophysiological estimates of the time course of semantic and
phonological encoding during listening and naming. Neuropsycho-
logia. 40:778--787.
Cerebral Cortex April 2010, V 20 N 4 927
by guest on Septem
ber 30, 2011cercor.oxfordjournals.org
Dow
nloaded from
Rugg MD, Doyle MC. 1994. Event-related potentials and stimulus repetition
in direct and indirect tests of memory. In: Heinze H, Munte T, Mangun
GR,editors.Cognitiveelectrophysiology.Boston:Birkhauser.p.124--148.
Salmelin R, Hari R, Lounasmaa OV, Sams M. 1994. Dynamics of brain
activation during picture naming. Nature. 368:463--465.
Sanchez-Casas R, Garcıa-Albea JE. 2005. The representation of cognate
and non cognate words on bilingual memory: can cognate status be
characterized as a special kind of morphological relation? In: Kroll J,
de Groot A, editors. Handbook of bilingualism: psycholinguistic
approaches. Oxford: Oxford University Press. p. 226--250.
Schiller NO, Bles M, Jansma BM. 2003a. Tracking the time course of
phonological encoding in speech production: an event-related brain
potential study. Brain Res. 17:819--831.
Schmitt BM, Munte TF, Kutas M. 2000. Electrophysiological estimates of
the time course of semantic and phonological encoding during
implicit picture naming. Psychophysiology. 37:473--484.
Schriefers H, Meyer AS, Levelt WJM. 1990. Exploring the time course of
lexical access in language-production: picture word interference
studies. J Mem Lang. 29:86--102.
Sebastian-Galles N, Martı M, Carreiras M, Cuetos F. 2000. LEXESP:
Lexico informatizado del espanol. Ediciones Universitat de Barce-
lona, Barcelona. [LEXESP: Spanish informatized lexic].
Sereno SC, Rayner K, Posner MI. 1998. Establishing a time-line of word
recognition: evidence from eye movement and event-related
potentials. Neuroreport. 9:2195--2200.
Sitnikova T, West WC, Kuperberg GR, Holcomb PJ. 2006. The neural
organization of semantic memory: electrophysiological activity
suggests feature-based segregation. Biol Psychol. 71:326--340.
Snodgrass JG, Vanderwart M. 1980. A standardized set of 260 pictures:
norm for name agreement, image agreement, familiarity, and visual
complexity. J Exp Psychol Hum Learn Mem. 6:174--215.
Thierry G, Cardebat D, Demonet JF. 2003. Electrophysiological
comparison of grammatical processing and semantic processing of
single spoken nouns. Brain Res. 17:535--547.
Thierry G, Doyon B, Demonet JF. 1998. ERP mapping in phonological
and lexical semantic monitoring tasks: a study complementing
previous PET results. Neuroimage. 8:391--408.
Thierry G, Martin CD, Downing P, Pegna AJ. 2007. Controlling for
interstimulus perceptual variance abolishes N170 face selectivity.
Nat Neurosci. 10:505--511.
Thorpe S, Fize D, Marlot C. 1996. Speed of processing in the human
visual system. Nature. 381:520--522.
Van Hell JG, De Groot AMB. 1998. Conceptual representation in
bilingual memory: effects of concreteness and cognate status in
word association. Bilingualism. 1:193--211.
Van Turennout M, Hagoort P, Brown CM. 1997. Electrophysiological
evidence on the time course of semantic and phonological
processes in speech production. J Exp Psychol Learn Mem Cogn.
23:787--806.
Van Turennout M, Hagoort P, Brown CM. 1998. Brain activity during
speaking: from syntax to phonology in 40 milliseconds. Science.
280:572--574.
Verhoef K, Roelofs A, Chwilla JC. 2009. Role of inhibition in language
switching: evidence from event-related brain potentials in overt
picture naming. Cognition. 110:84--99.
Wheeldon L, Levelt WJM. 1995. Monitoring the time course of
phonological encoding. J Mem Lang. 34:311--334.
Wohlert AB. 1993. Event-related brain potentials preceding speech and
nonspeech oral movements of varying complexity. Speech Hear Res.
36:905--987.
Wingfield A. 1968. Effects of frequency on identification and naming of
objects. Am J Psychol. 81:226--234.
928 The Time Course of Lexical Access in Speech Production d Strijkers et al.
by guest on Septem
ber 30, 2011cercor.oxfordjournals.org
Dow
nloaded from